Pedagogía de la educación a distancia

We show here that form3 is required for dendritic development, while the potential role of the human ortholog, INF2, is unknown with respect to neural development, however based on what is known regarding INF2 function, it is possible that there are evolutionarily conserved functions between these genes. For example, formation of stabilized MTs requires INF2, which was also found to be essential for centrosome reorientation in T cells (Laura Andrés-Delgado, 2012). The INF2 FH2 domain mediates the formation of stable, detyrosinated MTs in order to restore centrosome translocation (Laura Andrés-Delgado, 2012). Moreover, multiple studies have demonstrated that stable MTs plays an active role during the specification and promotion of neurite elaboration in early neuronal development (Witte et al. 2008, Falconer, 1989). In this study, we show that the morphological defects caused by disruption of form3 can be partially rescued by expression of the INF2 FH1-FH2 domains and that such INF2 expression, not only rescues dendritic morphology defects, but recovers dendritic MT stabilization. Therefore, our works defines a primordial role for form3/INF2 in regulation of dendritic architecture by regulating the MT cytoskeleton.

Mutations in INF2 are known to be causative for CMT dominant intermediate E disease (Boyer et al. 2011), although the mechanistic function of INF2 in disease pathogenesis are unclear. Neurological features of CMT include peripheral motor and sensory neuropathies, and the primary phenotypes consist of progressive distal muscle weakness and atrophy, reduced tendon reflexes, foot and hand deformities and peripheral insensitivity (Ekins et al. 2015). We observed that disruption of form3 in CIV nociceptive sensory neurons severely impairs behavioral responses to noxious heat, resulting in largely insensitive larval behavior, which is consistent with sensory

neuropathies observed in patients with CMT. Intriguingly, we can revert this heat insensitivity by the introduction of INF2 in the form3 mutant background revealing that there is not only morphological rescue, but also behavioral rescue. Previously, CMT diseases have been characterized by defects in axonal development, myelination, protein translation, and intracellular traffic of vesicles and organelles (Bucci et al. 2012; Niehues et al. 2014). Our work suggests an alternative mechanism that aberrant INF2 activity may impact: chiefly dendritic atrophy, as opposed to axonal defects, which leads to dramatically reduced dendritic field coverage manifesting as peripheral insensitivity to nociceptive thermal stimuli. Moreover, CMT disease has been linked to various defects in mitochondrial dynamics (Cassereau et al. 2011). CMT causing mutations have been shown to alter energy production via a mitochondrial complex I deficiency (Cassereau et al. 2011). We likewise observed defects in dendritic mitochondrial trafficking in

form3 mutants. Combined, our findings provide novel mechanistic insights into the potential etiological bases of INF2-mediated CMT sensory neuropathy, and provide evidence for functional conservation of these molecules between Drosophila and humans. Thus, we propose that

Drosophila da neurons represent a powerful platform for unraveling the mechanistic functions of these Formin molecules at both the morphological and behavioral levels, with direct implications for elucidating the neuronal bases of CMT sensory neuropathies.

REFERENCES CITED

Adryan, B., and Teichmann, S.A. 2010. “The Developmental Expression Dynamics of

Drosophila Melanogaster Transcription Factors.” Genome Biology 11 (4). BioMed Central Ltd: R40. doi:10.1186/gb-2010-11-4-r40.

Alexander, C., Votruba, M., Pesch, U.E., Thiselton, D.L., Mayer, S., Moore, A., Rodriguez, M., Kellner, U., Leo-Kottler, B., Auburger, G., Bhattacharya, S.S. and Wissinger B. 2000. “OPA1, encoding a dynamin-related GTPase, is mutated in autosomal dominant optic atrophy linked to chromosome 3q28.” Nat Genet. 26(2):211-5.

Andersen, E.F., S Asuri, N.S. and Halloran, M. C. 2011. “In Vivo Imaging of Cell Behaviors and F-Actin Reveals LIM-HD Transcription Factor Regulation of Peripheral versus Central Sensory Axon Development.” Neural Development 6 (1): 27. doi:10.1186/1749-8104-6-27. Andersen, R., Li, Y., Resseguie, M. and Brenman, J.E. 2005. “Calcium/calmodulin-Dependent

Protein Kinase II Alters Structural Plasticity and Cytoskeletal Dynamics in Drosophila.”

The Journal of Neuroscience : The Official Journal of the Society for Neuroscience 25 (39): 8878–88. doi:10.1523/JNEUROSCI.2005-05.2005.

Anderton, B.H., Callahan,L., Coleman. P., Davies, P., Flood, D., Jicha, G.A., Ohm, T. and Weaver, C. 1998. “Dendritic Changes in Alzheimer’s Disease and Factors That May Underlie These Changes.” Progress in Neurobiology 55 (6): 595–609.

Andrés-Delgado, L., Antón, O.M., Bartolini, F., Ruiz-Sáenz, A., Correas, I., Gundersen, G.G. and Alonso. M.A. 2012. “INF2 Promotes the Formation of Detyrosinated Microtubules Necessary for Centrosome Reorientation in T Cells.” The Journal of Cell Biology 198 (6): 1025–37. doi:10.1083/jcb.201202137.

Andrés-Delgado, L., Antón, O.M., Madrid, R., Byrne, J.A., and Alonso, M.A. 2010. “Formin INF2 Regulates MAL-Mediated Transport of Lck to the Plasma Membrane of Human T Lymphocytes.” Blood 116 (26): 5919–29. doi:10.1182/blood-2010-08-300665.

Armistead, J. and Triggs-Raine, B. (2014). Diverse diseases from a ubiquitous process: The ribosomopathy paradox. FEBS Letters 588: 1491-1500.

Arganda-Carreras, I., Fernández-González, R., Muñoz-Barrutia, A., Ortiz-De-Solorzano, C. (2010) 3D reconstruction of histological sections: Application to mammary gland tissue. Microsc Res Tech 73: 1019–1029doi:10.1002/jemt.20829

Baas, P. W., Deitch, J.S., Black, M.M. and Banker, G.A. 1988. “Polarity Orientation of

Microtubules in Hippocampal Neurons: Uniformity in the Axon and Nonuniformity in the Dendrite.” Proceedings of the National Academy of Sciences of the United States of America 85 (21): 8335–39.

Bartolini, F., and Gundersen, G.G. 2010. “Formins and Microtubules.” Biochimica et Biophysica Acta 1803 (2): 164–73. doi:10.1016/j.bbamcr.2009.07.006.

Barua, M., Brown, E.J., Charoonratana, V.T., Genovese, G., Sun, H. and Pollak, M.R. 2013. “Mutations in the INF2 Gene Account for a Significant Proportion of Familial but Not Sporadic Focal and Segmental Glomerulosclerosis.” Kidney International 83 (2): 316–22. doi:10.1038/ki.2012.349.

Baum, B., and Kunda, P. 2005. “Actin Nucleation: Spire - Actin Nucleator in a Class of Its Own.” Current Biology : CB 15 (8): R305–8. doi:10.1016/j.cub.2005.04.004.

Bellen, H.J., Tong, C. and Tsuda., H. 2010. “100 Years of Drosophila Research and Its Impact on Vertebrate Neuroscience: A History Lesson for the Future.” Nature Reviews

Neuroscience 11 (7): 514–22. doi:10.1038/nrn2839.

Belmonte, M.K., Allen, G., Beckel-Mitchener, A., Boulanger, L.M., Carper, R.A. and Webb, S.J. 2004. “Autism and Abnormal Development of Brain Connectivity.” The Journal of

Neuroscience : The Official Journal of the Society for Neuroscience 24 (42): 9228–31. doi:10.1523/JNEUROSCI.3340-04.2004.

Benjamini, Y. and Hochberg, Y. (1995). Controlling the false discovery rate: a practical and powerful approach to multiple testing. Journal of the Royal Statistical Society, Series B. 57 (1): 289–300. MR 1325392.

Benninger, Y., Thurnherr, T., Pereira, JA., Krause, S., Wu, X., Chrostek-Grashoff, A., Herzog, D., Nave, K.A., Franklin, R.J., Meijer, D., Brakebusch, C., Suter, U., and Relvas, J.B. (2007). Essential and Distinct Roles for cdc42 and rac1 in the Regulation of Schwann Cell Biology during Peripheral Nervous System Development. The Journal of Cell Biology 177 (6): 1051– 61. doi:10.1083/jcb.200610108.

Bhattacharya S, Iyer E.P., Iyer, S.C., Cox, D.N. (2014). Cell-type specific transcriptomic

profiling to dissect mechanisms of differential dendritogenesis. Genomics Data 2: 378-381. doi: 10.1016/j.gdata.2014.10.011

Bi, C., Wu, J., Jiang, T., Liu, Q., Cai, W., Yu, P., Cai, T., Zhao, M., Jiang, Y., Sun, Z. S. (2012). Mutations of ANK3 identified by exome sequencing are associated with autism

susceptibility. Hum. Mutat. 33: 1635-1638.

Blochlinger, K., Bodmer, R., Jack, J., Jan, L.Y. and Jan, Y.N. 1988. “Primary Structure and Expression of a Product from Cut, a Locus Involved in Specifying Sensory Organ Identity in Drosophila.” Nature 333 (6174): 629–35. doi:10.1038/333629a0.

Blochlinger, K., Bodmer, R., Jan, L.Y. and Jan, Y.N. 1990. “Patterns of Expression of Cut, a Protein Required for External Sensory Organ Development in Wild-Type and Cut Mutant Drosophila Embryos.” Genes & Development 4 (8): 1322–31.

Blochlinger, K., Jan, L.Y. and Jan, Y.N. 1993. “Postembryonic patterns of expression of cut, a locus regulating sensory organ identity in Drosophila.” Development. 117(2):441-50. Bodmer, R., Barbel, S., Sheperd, S., Jack, J.W., Jan, L.Y. and Jan, Y.N. 1987. “Transformation

of Sensory Organs by Mutations of the Cut Locus of D. Melanogaster.” Cell 51 (2): 293– 307.

Boyer, O., Nevo, F., Plaisier, E., Funalot, B., Gribouval, O., Benoit, G., Huynh Cong, E.,

Arrondel, C., Tête, M.J., Montjean, R., Richard, L., Karras, A., Pouteil-Noble, C., Balafrej, L., Bonnardeaux, A., Canaud, G., Charasse, C., Dantal, J., Deschenes, G., Deteix, P.,

Dubourg, O., Petiot, P., Pouthier, D., Leguern, E., Guiochon-Mantel, A., Broutin, I., Gubler, M.C., Saunier, S., Ronco, P., Vallat, JM., Alonso, M.A., Antignac, C., and Mollet, G. (2011). INF2 Mutations in Charcot-Marie-Tooth Disease with Glomerulopathy. The New England Journal of Medicine 365 (25): 2377–88. doi:10.1056/NEJMoa1109122.

Brand A.H. and Perrimon, N. 1993. Targeted gene expression as a menas of altering cell fates andgenerating dominant phenotypes. Development 118(2):401-15.

Breitsprecher, D. and Goode, B. L. 2013. “Formins at a Glance.” Journal of Cell Science 126 (Pt 1): 1–7. doi:10.1242/jcs.107250.

Brouhard, G. J., Stear, J. H., Noetzel, T. L., Al-Bassam, J., Kinoshita, K., Harrison, S. C., Howard, J., Hyman, A. A. (2008). XMAP215 is a processive microtubule polymerase. Cell

132: 79-88.

Brown, E.J., Schlöndorff, J.S., Becker, D.J., Tsukaguchi, H., Tonna, S.T., Uscinski, A.L., Higgs, H.N., Henderson, J.M. and Pollak, M.R. 2010. “Mutations in the Formin Gene INF2 Cause Focal Segmental Glomerulosclerosis.” Nature Genetics 42 (1): 72–76. doi:10.1038/ng.505. Bucci, C., Bakke, O., and Progida, C. (2012). Charcot-Marie-Tooth disease and intracellular

traffic. Prog Neurobiol 99:191-225.

Campellone, K.G. and Welch, M.D. 2010. “A Nucleator Arms Race: Cellular Control of Actin Assembly.” Nature Reviews. Molecular Cell Biology 11 (4): 237–51. doi:10.1038/nrm2867. Cannon, R.C., Turner, D.A., Pyapali, G.K. and Wheal, H. V. 1998. “An on-line archive of

reconstructed hippocampal neurons”. J. Neurosci. Methods, 84, 49–54.

Cassereau, J., Chevrollier, A., Gueguen, N., Desquiret, V., Verny, C., Nicolas, G., Dubas, F., Amati-Bonneau, P., Reynier, P., Bonneau, D., and Procaccio, V. (2011). Mitochondrial dysfunction and pathophysiology of Charcot-Marie-Tooth disease involving GDAP1

mutations. Exp Neurology 227:31-41.

Castón J.R. (2013) Conventional electron microscopy, cryo-electron microscopy and cryo- electron tomography of viruses. Subcell Biochem 68: 79-115. doi:10.1007/978-94-007-

6552-8_3

Chan, D.C. 2006. “Mitochondrial fusion and fission in mammals.” Annu Rev Cell Dev Biol.

22:79-99.

Chang, F. 2000. “Microtubule and Actin-Dependent Movement of the Formin cdc12p in Fission Yeast.” Microscopy Research and Technique 49 (2): 161–67. doi:10.1002/(SICI)1097- 0029(20000415)49:2<161::AID-JEMT8>3.0.CO;2-2.

Chen, F., Archambault, V., Kar, A., Lio, P., D'Avino, P.P., Sinka, R., Lilley, K., Laue, E.D., Deak, P., Capalbo, L., Glover, D.M. (2007). Multiple protein phosphatases are required for mitosis in Drosophila. Curr. Biol. 17(4): 293-303.

Chesarone, M.A., DuPage, A.G. and Goode, B.L. 2010. “Unleashing Formins to Remodel the Actin and Microtubule Cytoskeletons.” Nature Reviews. Molecular Cell Biology 11 (1): 62– 74. doi:10.1038/nrm2816.

Chhabra, E.S., and Higgs, H.N. 2007. “The Many Faces of Actin: Matching Assembly Factors with Cellular Structures.” Nature Cell Biology 9 (10): 1110–21. doi:10.1038/ncb1007-1110. Chiang, A.S., Lin, C.Y., Chuang, C.C., Chang, H.M., Hsieh, C.H., Yeh, C.W., Shih, C.T., Wu,

J.J., Wang, G.T., Chen, Y.C., Wu, C.C., Chen, G.Y., Ching, Y.T., Lee, P.C., Lin, C.Y., Lin, H.H., Wu, C.C., Hsu, H.W., Huang, Y.A., Chen, J.Y., Chiang, H.J., Lu, C.F., Ni, R.F., Yeh. C.Y. and Hwang J.K. 2011. “Three-Dimensional Reconstruction of Brain-Wide Wiring Networks in Drosophila at Single-Cell Resolution.” Current Biology : CB 21 (1): 1–11. doi:10.1016/j.cub.2010.11.056.

Cline, H.T. 2001. “Dendritic Arbor Development and Synaptogenesis.” Current Opinion in Neurobiology 11 (1): 118–26. doi:10.1016/S0959-4388(00)00182-3.

Coles, C.H., and Bradke, F. 2015. “Coordinating Neuronal Actin – Microtubule Dynamics.”

Current Biology 25 (15). Elsevier Ltd: R677–91. doi:10.1016/j.cub.2015.06.020.

Conde, C., and Cáceres, A. 2009. “Microtubule Assembly, Organization and Dynamics in Axons and Dendrites.” Nature Reviews. Neuroscience 10 (5). Nature Publishing Group: 319–32. doi:10.1038/nrn2631.

Corty, M.M., Matthews, B.J. and Grueber,W.B. 2009. “Molecules and Mechanisms of Dendrite Development in Drosophila.” Development (Cambridge, England) 136 (7): 1049–61. doi:10.1242/dev.014423.

Couton, L., Mauss, A.S., Yunusov, T., Diegelmann, S., Evers, J.F. and Landgraf, M. 2015. “Development of Connectivity in a Motoneuronal Network in Drosophila Larvae.” Current Biology 25 (5): 568–76. doi:10.1016/j.cub.2014.12.056.

Crowner, D., Madden, K., Goeke, S. and Giniger, E. 2002. “Lola Regulates Midline Crossing of CNS Axons in Drosophila.” Development (Cambridge, England) 129 (6): 1317–25.

Crozatier, M., and Vincent, A. 2008. “Control of Multidendritic Neuron Differentiation in Drosophila: The Role of Collier.” Developmental Biology 315 (1): 232–42.

doi:10.1016/j.ydbio.2007.12.030.

Cubelos, B., Sebastián-Serrano, A., Beccari, L., Calcagnotto, M.E., Cisneros, E., Kim, S., Ana Dopazo, A., Alvarez-Dolado, M., Redondo, J.M., Bovolenta, P., Walsh, C.A., Nieto, M. 2010. “Cux1 and Cux2 Regulate Dendritic Branching, Spine Morphology, and Synapses of the Upper Layer Neurons of the Cortex.” Neuron 66 (4): 523–35.

doi:10.1016/j.neuron.2010.04.038.

Cuntz, H., Forstner, F., Borst, A. and Häusser, M. 2010. “One rule to grow them all: A general theory of neuronal branching and its practical application.” PLoS Comput. Biol., 6

Cvrčková, Fa. 2013. “Formins and Membranes: Anchoring Cortical Actin to the Cell Wall and Beyond.” Frontiers in Plant Science 4 (January): 436. doi:10.3389/fpls.2013.00436. De Rechter, S., Waele, L.D., Levtchenko, E. and Mekahli. D. 2015. “Charcot-Marie-Tooth: Are

You Testing for Proteinuria?” European Journal of Paediatric Neurology : EJPN : Official Journal of the European Paediatric Neurology Society 19 (1): 1–5.

doi:10.1016/j.ejpn.2014.08.004.

de la Torre-Ubieta, L., and Bonni, A. 2011. “Transcriptional Regulation of Neuronal Polarity and Morphogenesis in the Mammalian Brain.” Neuron 72 (1): 22–40.

doi:10.1016/j.neuron.2011.09.018.

de Taffin, M., Carrier, Y., Dubois, L., Bataillé, L., Painset, A., Le Gras, S., Jost, B., Crozatier, M., Vincent, A. (2015). Genome-Wide Mapping of Collier In Vivo Binding Sites Highlights Its Hierarchical Position in Different Transcription Regulatory Networks. PloS One, 10(7), e0133387.

Deciphering Developmental Disorders Study. (2015). Large-scale discovery of novel genetic causes of developmental disorders. Nature 519: 223-228.

Deeks, M.J, Fendrych, M., Smertenko, A., Bell, K.S., Oparka, K., Cvrcková, F., Zársky, V. and Hussey, P.J. 2010. “The Plant Formin AtFH4 Interacts with Both Actin and Microtubules, and Contains a Newly Identified Microtubule-Binding Domain.” Journal of Cell Science

123 (Pt 8): 1209–15. doi:10.1242/jcs.065557.

Dehmelt, L. and Halpain, S. 2004. “Actin and Microtubules in Neurite Initiation: Are MAPs the Missing Link?” Journal of Neurobiology 58 (1): 18–33. doi:10.1002/neu.10284.

Deitch, J.S., and Rubel, E.W. 1984. “Afferent Influences on Brain Stem Auditory Nuclei of the Chicken: Time Course and Specificity of Dendritic Atrophy Following Deafferentation.”

The Journal of Comparative Neurology 229 (1): 66–79. doi:10.1002/cne.902290106. Denk, W., Strickler, J.H. and Webb, W.W. (1990). Two-photon laser scanning fluorescence

microscopy. Science 248:73–76. doi:10.1126/science.2321027

Denk W. and Svoboda, K. (1997). Photon upmanship: why multiphoton imaging is more than a gimmick. Neuron 18:351–357. doi:10.1002/cne.902290106

D’Este E., Kamin, D., Gottfert, F., El-Hady, A., Hell, S.W. (2015). STED nanoscopy reveals the ubiquity of subcortical cytoskeleton periodicity in living neurons. Cell Rep 10:1246-51. Detmer, S.A. and Chan, D.C. 2007. “Functions and dysfunctions of mitochondrial dynamics.”

Nat Rev Mol Cell Biol. 8(11):870-9.

Dickson, D.W., , Liu, W., Hardy, J., Farrer, M ., Mehta, N., Uitti, R., Mark, M., Zimmerman, T., Golbe, L., Sage, J., Sima, A., D'Amato, C., Albin, R., Gilman, S., Yen, S.H. 1999. “Widespread Alterations of Alpha-Synuclein in Multiple System Atrophy.” The American Journal of Pathology 155 (4): 1241–51.

DiVincenzo, C., Elzinga, C.D., Medeiros, A.C., Karvassi, I., Jones, J.R., Evans, M.C., Braastad, C.D., Bishop, C.M., Jaremko, M., Wang, Z., Liaquat, K., Hoffman, C.A., York, M.D., Batish, S.D., Lupski, J.R. and Higgins, J.J. (2014). The allelic spectrum of Charcot-Marie- Tooth disease in over 17,000 individuals with neuropathy. Mol Genet Genomic Med

2(6):522-529.

Dollar, G., Gombos, R., Barnett, A.A., Hernandez, D.S., Maung, S.M., Mihaly, J., and Jenny, A. (2016). “Unique and overlapping functions of Formins Frl and DAAM during ommatidial rotation and neuronal development in Drosophila.” Genetics 202:1135-1151.

Dunn, AY, Melville MW, and Frydman J (2001). Review: cellular substrates of the eukaryotic chaperonin TRiC/CCT. J Struct Biol 135(2): 176-184.

Ekins, S., Kitterman, N.K., Arnold, R.J.G., Burgess, R.W., Freundlich, J.S., Gray, S.J., Higgins, J.J., Langley, B., Willis, D.E., Notterpek, L., Pleasure, D., Sereda, M.W., and Moore, A. (2015). A brief review of recent Charcot-Marie-Tooth research and priorities.

F1000Research 4:53, doi: 10.12688/f1000research.6160.1.

Evangelista, M., Pruyne, D., Amberg, D. C., Boone, C. and Bretscher, A. 2002. “Formins direct Arp2/3-independent actin filament assembly to polarize cell growth in yeast.” Nat. Cell Biol. 4, 32 -41.

Falconer, M.M., Vielkind, U. and Brown, D.L. 1989. “Establishment of a stable, acetylated microtubule bundle during neuronal commitment.” Cell Motil Cytoskeleton.12(3):169-80. Feng, L., Zhao, T., and Kim, J. 2015. “neuTube 1.0: A New Design for Efficient Neuron

Reconstruction Software Based on the SWC Format.” eNeuro, 2, ENEURO.0049-14.2014. Ferreira, T., Ou, Y., Li, S., Giniger, E. and Meyel, D.J. 2014. “Dendrite Architecture Organized

by Transcriptional Control of the F-Actin Nucleator Spire.” Development (Cambridge, England) 141 (3): 650–60. doi:10.1242/dev.099655.

Fiala, J.C., Spacek, J. and Harris, K.M. 2002. “Dendritic Spine Pathology: Cause or

Consequence of Neurological Disorders?” Brain Research. Brain Research Reviews 39 (1): 29–54.

Förster, D., Armbruster, K. and Luschnig, S. (2010). Sec24-dependent secretion drives cell autonomous expansion of tracheal tubes in Drosophila. Curr Biol 20:62-68.

Frank, M. 2000. “MAL, a Proteolipid in Glycosphingolipid Enriched Domains: Functional Implications in Myelin and Beyond.” Progress in Neurobiology 60 (6): 531–44. Franker, M.A.M, and Hoogenraad, C.C. 2013. “Microtubule-Based Transport - Basic

Mechanisms, Traffic Rules and Role in Neurological Pathogenesis.” Journal of Cell Science

126 (Pt 11): 2319–29. doi:10.1242/jcs.115030.

Friedman, J.R., Lackner, L.L., West, M., DiBenedetto, J.R., Nunnari, J. and Voeltz, G.K. 2011. “ER tubules mark sites of mitochondrial division.” Science. 334(6054):358-62. doi: 10.1126/science.1207385.

Gaillard, J., Ramabhadran, V., Neumanne, E., Gurel, P., Blanchoin, L., Vantard, M. and Higgs, H.N. 2011. “Differential Interactions of the Formins INF2, mDia1, and mDia2 with Microtubules.” Molecular Biology of the Cell 22 (23): 4575–87. doi:10.1091/mbc.E11-07- 0616.

Gao, F.B., Brenman, J.E., Jan, L.Y. and Jan, Y.N. 1999. “Genes Regulating Dendritic

Outgrowth, Branching, and Routing in Drosophila.” Genes & Development 13 (19): 2549– 61.

Garey, L.J., Ong, W.Y., Patel, T.S., Kanani, M., Davis, A., Mortimer, A.M., Barnes, T.R. and R H.S. 1998. “Reduced Dendritic Spine Density on Cerebral Cortical Pyramidal Neurons in Schizophrenia.” Journal of Neurology, Neurosurgery, and Psychiatry 65 (4): 446–53. Gasteier, J.E., Schroeder, S., Muranyi, W., Madrid, R., Benichou, S. and Fackler, O.T. 2005.

“FHOD1 Coordinates Actin Filament and Microtubule Alignment to Mediate Cell Elongation.” Experimental Cell Research 306 (1): 192–202.

doi:10.1016/j.yexcr.2005.02.006.

Gates, M., Kannan, R. and Giniger E. 2011. “A Genome-Wide Analysis Reveals That the Drosophila Transcription Factor Lola Promotes Axon Growth in Part by Suppressing Expression of the Actin Nucleation Factor Spire.” Neural Development 6 (1). BioMed Central Ltd: 37. doi:10.1186/1749-8104-6-37.

Ghosh-Roy, A., Desai, B.S., Ray, K. (2005). Dynein light chain 1 regulates dynamin-mediated F-actin assembly during sperm individualization in Drosophila. Mol. Biol. Cell 16(7): 3107-3116.

Giniger, E., Tietje, K., Jan, L.Y., and Jan, Y.N. 1994. “Lola Encodes a Putative Transcription Factor Required for Axon Growth and Guidance in Drosophila.” Development (Cambridge, England) 120 (6): 1385–98.

Goley, E.D. and Welch, M.D. 2006. “The ARP2/3 complex: an actin nucleator comes of age.”

Nat Rev Mol Cell Biol. 2006 Oct;7(10):713-26.

Goode, B.L., and Eck M.J. 2007. “Mechanism and Function of Formins in the Control of Actin